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The Entotympanic of Pangolins and the Phylogeny of the Pholidota (Mammalia)

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Abstract

Entotympanics are independent elements present in the auditory bullae of various eutherians. An entotympanic has been reported for extant pangolins of the Order Pholidota, but the actual distribution of this element remains uncertain, in part, because it is a small, loosely attached structure that is often lost in macerated skulls. Consequently, it is unknown whether or not the entotympanic characterizes Pholidota primitively or has evolved within the group. This report addresses the morphology and distribution of the entotympanic among living and extinct pholidotans. An entotympanic occurs in the African pangolins Manis gigantea, M. temminckii, and in one specimen of M. tricuspis. In each, it is a small, nodular bone that occupies a distinct fossa primarily on the basioccipital, the presence of which allows us to assess the occurrence of an entotympanic even in specimens in which the bone has fallen out. Both the entotympanic and the basioccipital facet are lacking in the four remaining extant pangolin species and in the late Eocene pangolin Patriomanis. To assess the significance of this entotympanic distribution, a phylogenetic analysis of extant pangolins plus Patriomanis based on 67 cranial characters was performed. Four different outgroup analyses all resulted in the same single most parsimonious tree, in which the three extant Asian pangolins form a monophyletic clade and the four extant African pangolins fall into a paraphyletic assemblage. Optimization of the entotympanic distribution onto this tree results in two patterns, dependent on the outgroup choice. If Patriomanis is the sole outgroup to the extant pangolins, the entotympanic arises within pangolins as a synapomorphy of Manis gigantea and M. temminckii, convergently acquired in some M. tricuspis. If Xenarthra and Palaeanodonta are employed as outgroups, the entotympanic optimization is ambiguous: the pattern is either as above or the entotympanic is present primitively within Pholidota and lost secondarily in Patriomanis and a clade comprising M. tricuspis, M. tetradactyla, and the Asian forms.

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LITERATURE CITED

  • Barlow, J. C. (1984). Xenarthrans and pholidotes. In: Orders and Families of Recent Mammals of the World, S. Anderson and J. Knox Jones, Jr., eds., pp. 219–239, John Wiley & Sons, New York.

    Google Scholar 

  • Cifelli, R. L. (1983). Eutherian tarsals from the late Paleocene of Brazil. Am. Mus. Novitates 2761: 1–31.

    Google Scholar 

  • Corbet, G. B., and Hill, J. E. (1991). A World List of Mammalian Species, 3rd ed., British Museum (Natural History), London.

    Google Scholar 

  • Czelusniak, J., Goodman, M., Koop, B. F., Tagle, D. A., Shoshani, J., Braunitzer, G., Kleinschmidt, T. K., DeJong, W. W., and Matsuda, G. (1990). Perspectives from amino acid and nucleotide sequences on cladistic relationships among higher taxa of Eutheria. In: Current Mammalogy, Vol. 2, H. H. Genoways, ed., pp. 545–572, Plenum Press, New York.

    Google Scholar 

  • De Beer, G. R. (1937). The Development of the Vertebrate Skull, Clarendon Press, Oxford.

    Google Scholar 

  • Emry, R. J. (1970). A North American Oligocene pangolin and other additions to the Pholidota. Bull. Am. Mus. Nat. Hist. 142: 455–510.

    Google Scholar 

  • Engelmann, G. F. (1978). The Logic of Phylogenetic Analysis and the Phylogeny of the Xenarthra (Mammalia), Ph.D. dissertation, Columbia University, New York.

    Google Scholar 

  • Engelmann, G. F. (1985). The phylogeny of the Xenarthra. In: The Ecology and Evolution of Armadillos, Sloths, and Vermilinguas, G. G. Montgomery, ed., pp. 51–64, Smithsonian Institution Press, Washington, DC.

    Google Scholar 

  • Eschweiler, R. (1899). Zur vergleichenden Anatomie der Muskeln und der Topographie des Mittelohres verschiedener Säugethiere. Arch. Mikr. Anat. 53: 558–622.

    Google Scholar 

  • Frechkop, S. (1931). Notes sur les mammifères. VI. Quelques observations sur la classification des pangolins (Manidae). Bull. Mus. Roy. Sci. Nat. Belg. 7: 1–14.

    Google Scholar 

  • Gaudin, T. J. (1993). Phylogeny of the Tardigrada (Mammalia, Xenarthra) and the Evolution of Locomotor Function in the Xenarthra, Ph.D. dissertation, University of Chicago, Chicago.

    Google Scholar 

  • Gaudin, T. J. (1995). The ear region of edentates and the phylogeny of the Tardigrada (Mammalia, Xenarthra). J. Vert. Paleontol. 15: 672–705.

    Google Scholar 

  • Gaudin, T. J., and Branham, D. G. (1998). The phylogeny of the Myrmecophagidae (Mammalia, Xenarthra, Vermilingua) and the relationship of Eurotamandua to the Vermilingua. J. Mammal. Evol. 5: 237–265.

    Google Scholar 

  • Gaudin, T. J., Wible, J. R., Hopson, J. A., and Turnbull, W. D. (1996). Reexamination of the morphological evidence for the Cohort Epitheria (Mammalia, Eutheria). J. Mammal. Evol. 3: 31–79.

    Google Scholar 

  • Glass, B. P. (1985). History of classification and nomenclature in Xenarthra (Edentata). In: The Ecology and Evolution of Armadillos, Sloths, and Vermilinguas, G. G. Montgomery, ed., pp. 1–3, Smithsonian Institution Press, Washington, DC.

    Google Scholar 

  • Grassé, P.-P. (1955). Ordre de Pholidotes. In: Traité de Zoologie, Vol. 17, Mammifères, P.-P. Grassé, ed., pp. 1267–1282, Masson et Cie, Paris.

    Google Scholar 

  • Graur, D., Duret, L., and Gouy, M. (1996). Phylogenetic position of the order Lagomorpha (rabbits, hares and allies). Nature 379: 333–335.

    PubMed  Google Scholar 

  • Hershkovitz, P. (1992). Ankle bones: The Chilean opposum Dromiciops gliroides Thomas, and marsupial phylogeny. Bonn. Zool. Beitr. 43: 181–213.

    Google Scholar 

  • Hoffstetter, R. (1982). Les édentés xenartres, un groupe singulier de la faune neotropical. In: Paleontology, Essential of Historical Geology, E. M. Gallitelli, ed., pp. 385–443, STEM Mocchi Modena Press, Modena, Italy.

    Google Scholar 

  • Jollie, M. (1968). The head skeleton of a new-born Manis javanica with comments on the ontogeny and phylogeny of the mammal head skeleton. Acta Zool. 49: 227–305.

    Google Scholar 

  • Koenigswald, W. v., and Martin, T. (1990). Ein Skelett von Necromanis franconica, einem Schuppentier (Pholidota, Mammalia) aus dem Aquitan von Saulcet im Allier-Becken (Frankreich). Ecol. Geol. Helv. 83: 845–864.

    Google Scholar 

  • MacPhee, R. D. E. (1979). Entotympanics, ontogeny and primates. Folia Primatol. 31: 23–47.

    PubMed  Google Scholar 

  • MacPhee, R. D. E. (1981). Auditory regions of primates and eutherian insectivores: Morphology, ontogeny and character analysis. Contrib. Primatol. 18: 1–282.

    Google Scholar 

  • MacPhee, R. D. E. (1994). Morphology, adaptations, and relationships of Plesiorycteropus, and a diagnosis of a new order of eutherian mammals. Bull. Am. Mus. Nat. Hist. 220: 1–214.

    Google Scholar 

  • MacPhee, R. D. E., and Novacek, M. J. (1993). Definition and relationships of Lipotyphla. In: Mammal Phylogeny: Placentals, F. S. Szalay, M. J. Novacek, and M. C. McKenna, eds., pp. 13–31, Springer-Verlag, New York.

    Google Scholar 

  • Maier, W. (1989). Morphologische Untersuchungen am Mittelohr der Marsupialia. Z. Zool. Syst. Evol.-Forsch. 27: 149–168.

    Google Scholar 

  • Matthew, W. D. (1918). A revision of the lower Eocene Wasatch and Wind River faunas. Part V. Insectivora (continued), Glires, Edentata. Bull. Am. Mus. Nat. Hist. 38: 565–657.

    Google Scholar 

  • McKenna, M. C. (1992). The alpha crystallin A chain of the eye lens and mammalian phylogeny. Ann. Zool. Fenn. 28: 349–360.

    Google Scholar 

  • McKenna, M. C., and Bell, S. K. (1997). Classification of Mammals Above the Species Level, Colombia University Press, New York.

    Google Scholar 

  • Miyamoto, M. M., and Goodman, M. (1986). Biomolecular systematics of eutherian mammals: Phylogenetic pattern and classification. Syst. Zool. 35: 230–240.

    Google Scholar 

  • Norman, J. E., and Ashley, M. V. (1994). Molecular systematics of the xenarthrans. Am. Soc. Mamm. 75th Anniv. Mtg., Abstr. 271.

  • Norris, C. A. (1993). Changes in the composition of the auditory bulla in southern Solomon Islands populations of the grey cuscus, Phalanger orientalis breviceps (Marsupialia, Phalangeridae). Zool. J. Linn. Soc. 107: 93–106.

    Google Scholar 

  • Novacek, M. J. (1977). Aspects of the problem of variation, origin and evolution of the eutherian auditory bulla. Mammal Rev. 7: 131–149.

    Google Scholar 

  • Novacek, M. J. (1986). The skull of leptictid insectivorans and the higher-level classification of eutherian mammals. Bull. Am. Mus. Nat. Hist. 183: 1–112.

    Google Scholar 

  • Novacek, M. J. (1992). Mammalian phylogeny: Shaking the tree. Nature 356: 121–125.

    PubMed  Google Scholar 

  • Novacek, M. J. (1993). Patterns of diversity in the mammalian skull. In: The Skull, Vol. 2: Patterns of Structural and Systematic Diversity, J. Hanken and B. K. Hall, eds., pp. 438–545, University of Chicago Press, Chicago.

    Google Scholar 

  • Novacek, M. J., and Wyss, A. (1986). Higher-level relationships of the recent eutherian orders: morphological evidence. Cladistics 2: 257–287.

    Google Scholar 

  • Novacek, M. J., Wyss, A. R., and McKenna, M. C. (1988). The major groups of eutherian mammals. In: The Phylogeny and Classification of Tetrapods, Vol. 2. Mammals, M. J. Benton, ed., pp. 31–71, Clarendon Press, Oxford.

    Google Scholar 

  • Nowak, R. M. (1991). Walker's Mammals of the World, 5th ed., Johns Hopkins University Press, Baltimore.

    Google Scholar 

  • Patterson, B. (1978). Pholidota and Tubulidentata. In: Evolution of African Mammals, V. J. Maglio and H. B. S. Cooke, eds., pp. 268–278, Harvard University Press, Cambridge, MA.

    Google Scholar 

  • Patterson, B., Segall, W., and Turnbull, W. D. (1989). The ear region in xenarthrans (=Edentata, Mammalia). Part I. Cingulates. Fieldiana Geol. n.s. 18: 1–46.

    Google Scholar 

  • Patterson, B., Segall, W., Turnbull, W. D., and Gaudin, T. J. (1992). The ear region in xenarthrans (=Edentata, Mammalia). Part II. Pilosa (sloths, anteaters), palaeanodonts, and a miscellany. Fieldiana Geol. n.s. 24: 1–79.

    Google Scholar 

  • Pocock, R. I. (1924). The external characters of the pangolins (Manidae). Proc. Zool. Soc. London 1924: 707–723.

    Google Scholar 

  • Reiss, K. Z. (1996). Evolution of the feeding apparatus in ant-eating mammals: When do we stop believing in convergence? Am. Zool. 36(5): 85A.

    Google Scholar 

  • Reiss, K. Z. (1997). Myology of the feeding apparatus of myrmecophagid anteaters (Xenarthra: Myrmecophagidae). J. Mammal. Evol. 4: 87–117.

    Google Scholar 

  • Rose, K. D. (1979). A new Paleocene palaeanodont and the origin of the Metacheiromyidae (Mammalia). Breviora Mus. Comp. Zool. 455: 1–14.

    Google Scholar 

  • Rose, K. D., and Emry, R. J. (1993). Relationships of Xenarthra, Pholidota, and fossil “edentates.” In: Mammal Phylogeny: Placentals, F. S. Szalay, M. J. Novacek, and M. C. McKenna, eds., pp. 81–102, Springer-Verlag, New York.

    Google Scholar 

  • Sarich, V. M. (1985). Xenarthran systematics: albumin immunological evidence. In: The Evolution and Ecology of Armadillos, Sloths, and Vermilinguas, G. G. Montgomery, ed., pp. 77–81, Smithsonian Institution Press, Washington, DC.

    Google Scholar 

  • Schlitter, D. A. (1993). Order Pholidota. In: Mammal Species of the World, D. E. Wilson and D. M. Reeder, eds., p. 415, Smithsonian Institution Press, Washington, DC.

    Google Scholar 

  • Schneider, R. (1955). Zur Entwicklung des Chondrocraniums der Gattung Bradypus. Gegenbaurs Morph. Jb. 95: 209–301.

    Google Scholar 

  • Segall, W. (1973). Characteristics of the ear, especially the middle ear in fossorial mammals, compared with those in the Manidae. Acta Anat. 86: 96–110.

    PubMed  Google Scholar 

  • Simpson, G. G. (1931). Metacheiromys and the relationships of the Edentata. Bull. Am. Mus. Nat. Hist. 59: 295–381.

    Google Scholar 

  • Simpson, G. G. (1945). The principles of classification and a classification of mammals. Bull. Am. Mus. Nat. Hist. 85: 1–350.

    Google Scholar 

  • Spatz, W. B. (1966). Zur Ontogenese der Bulla tympanica von Tupaia glis Diard 1820 (Prosimiae, Tupaiiformes). Folia Primatol. 4: 26–50.

    PubMed  Google Scholar 

  • Springer, M. S., Cleven, G. C., Madsen, O., de Jong, W. W., Waddell, V. G., Amrine, H. M., and Stanhope, M. J. (1997). Endemic African mammals shake the phylogenetic tree. Nature 388: 61–64.

    PubMed  Google Scholar 

  • Starck, D. (1941). Zur Morphologie des Primordialkraniums von Manis javanica Desm. Gegenbaurs Morph. Jb. 86: 1–122.

    Google Scholar 

  • Storch, G. (1978). Eomanis waldi, ein Schuppentier aus dem Mittel-Eozän der “Grube Messel” bei Darmstadt (Mammalia: Pholidota). Senckenberg. lethaea 59: 503–529.

    Google Scholar 

  • Storch, G., and Martin, T. (1994). Ecomanis krebsi, ein neues Schuppentier aus dem Mittel-Eozän der Grube Messel bei Darmstadt (Mammalia: Pholidota). Berliner geowiss. Abh. E13: 83–97.

    Google Scholar 

  • Swofford, D. L. (1993). PAUP: Phylogenetic Analysis Using Parsimony, Version 3.1.1, Smithsonian Institution, Washington, DC.

    Google Scholar 

  • Szalay, F. S. (1977). Phylogenetic relationships and a classification of eutherian Mammalia. In: Major Patterns in Vertebrate Evolution, M. K. Hecht, P. C. Goody, and B. M. Hecht, eds., pp. 315–374, Plenum Press, New York.

    Google Scholar 

  • van Kampen, P. N. (1905). Die Tympanalgegend des Säugetiereschädels. Gegenbaurs Morph. Jb. 34: 321–722.

    Google Scholar 

  • van Kampen, P. N. (1915). De Phylogenie van het entotympanicum. Tijdschr. Ned. Dierkd. Ver. 14: xxiv.

    Google Scholar 

  • van der Klaauw, J. C. (1922). Über die Entwicklung des Entotympanicums. Tijdschr. Ned. Dierkd. Ver. 18: 135–174.

    Google Scholar 

  • van der Klaauw, J. C. (1931). On the auditory bulla in some fossil mammals. Bull. Am. Mus. Nat. Hist. 62: 1–352.

    Google Scholar 

  • Wible, J. R. (1984). The Ontogeny and Phylogeny of the Mammalian Cranial Arterial Pattern, Ph.D. dissertation, Duke University, Durham, NC.

    Google Scholar 

  • Wible, J. R. (1986). Transformations in the extracranial course of the internal carotid artery in mammalian phylogeny. J. Vert. Paleontol. 6: 313–325.

    Google Scholar 

  • Wible, J. R., and Martin, J. R. (1993). Ontogeny of the tympanic floor and roof in archontans. In: Primates and Their Relatives in Phylogenetic Perspective, R. D. E. MacPhee, ed., pp. 111–148, Plenum Press, New York.

    Google Scholar 

  • Wible, J. R., and Novacek, M. J. (1988). Cranial evidence for the monophyletic origin of bats. Am. Mus. Novitates 2911: 1–19.

    Google Scholar 

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Authors and Affiliations

  1. Department of Biological and Environmental Sciences, University of Tennessee at Chattanooga, 615 McCallie Avenue, Chattanooga, Tennessee, 37403-2598

    Timothy J. Gaudin

  2. Section of Mammals, Carnegie Museum of Natural History, 5800 Baum Boulevard, Pittsburgh, Pennsylvania, 15206-3706

    John R. Wible

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Gaudin, T.J., Wible, J.R. The Entotympanic of Pangolins and the Phylogeny of the Pholidota (Mammalia). Journal of Mammalian Evolution 6, 39–65 (1999). https://doi.org/10.1023/A:1020538313412

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